Invasive exotic species threaten native biodiversity, alter ecosystem structure and function, and annually cost over $100 billion in the US alone. Determining the ecological traits and interactions that affect invasion success are thus critical for predicting, preventing, and mitigating the negative effects of biological invasions. Native herbivores are widely assumed to facilitate exotic plant invasions by preferentially consuming native plants and avoiding exotic plants. Here, I use freshwater plant communities scattered broadly across the Southeastern U.S. to show that herbivory is an important force driving the ecology and evolution of freshwater systems. However, native consumers often preferentially consume rather than avoid exotic over native plants. Analyses of 3 terrestrial datasets showed similar patterns, with native herbivores generally preferring exotic plants. Thus, exotic plants appear defensively nave against these evolutionarily novel consumers, and exotic plants may escape their coevolved, specialist herbivores only to be preferentially consumed by the native generalist herbivores in their new ranges. In further support of this hypothesis, a meta-analysis of 71 manipulative field studies including over 100 exotic plant species and 400 native plant species from terrestrial, aquatic, and marine systems revealed that native herbivores strongly suppressed exotic plants, while exotic herbivores enhanced the abundance and species richness of exotic plants by suppressing native plants. Both outcomes are consistent with the hypothesis that prey are susceptible to evolutionarily novel consumers. Thus, native herbivores provide biotic resistance to plant invasions, but the widespread replacement of native with exotic herbivores eliminates this ecosystem service, facilitates plant invasions, and triggers an invasional meltdown. Consequently, rather than thriving because they escape their co-evolved specialist herbivores, exotic plants may thrive because their co-evolved generalist herbivores have stronger negative effects on evolutionarily nave, native plants.

<p>In the scope of future population support, agricultural productivity, in particular in sub-Saharan Africa, has to increase drastically to meet the UN’s millennium development goals of eradicating extreme poverty and hunger by 2015. Water availability in the root-zone limits crop production in large parts of the developing world. As competition for fresh water increases, water of lower quality, for example saline or polluted water, is often used for irrigation. Low-cost drip systems are suitable for saline water irrigation because they effectuate a minimisation of salt accumulation, leaf burn and peaks in salt concentration. Nonetheless, all types of saline water irrigation contain the risk for causing soil salinisation. Thus, in order to achieve long-term sustainability of these systems, appropriate management strategies are needed. The choice of management practices may be influenced by local conditions such as climate, soil and irrigation water salinity. A litera-ture review showed that there is a potential for saline water irrigation in sub-Saharan Africa in water scarce areas. Low-cost drip irrigation with saline water (6 dS m-1) was successfully used to irrigate two consecutive crops of tomato in semi-arid South Africa. An integrated ecosystems model was developed to simulate long-term yield and salt accumulation in a drip-irrigated agricultural system for a range of salinities, climates and management techniques. Crop, salt and water balance data from two field experiments conducted in Israel and South Africa, respectively, were used to parameterise and test the model. Emphasis was placed on testing the usability of the model as a tool for evaluating the importance of certain plausible management options of low-cost, drip-irrigation systems. Therefore, particular focus was directed towards correctly describing soil salinity stress on plant growth and soil evaporation from a distributed (wetted and dry) surface. In addition, the model was developed to function for different climates without having to change any other parameters or variables except for the actual climatic data. Simulations were subsequently run over a 30-year period to study long-term yield and salt accumulation in the soil profile for two sites in South Africa, demonstrating the applicability of the model. Model simulations showed that high soil salinities reduced crop growth and thus increased both drainage and soil evaporation. Further, covering the soil with a plastic sheet led to a reduction of soil evaporation and a subsequent increase in both transpiration and drainage. Rainfall was crucial for the leaching of salts from the soil, and thus in regions with low levels of rainfall, a higher leaching fraction of supplied saline irrigation water has to compensate for the lack of rain. However, a high leaching fraction also causes large amounts of salt leaching, which could potentially pollute underlying groundwater and downstream ecosystems. This risk can be mitigated using mulching, which minimises non-productive water losses, thereby lowering irrigation water needs. The choice of irrigation water salinity, frequency of irrigation and soil coverage may differ between the farmer and the regional water manager due to different preferences. Furthermore, the study highlighted how environmental variables such as water use efficiency and radiation use efficiency can be used as indicators of system performance. Whereas the latter is first and foremost a general stress indicator, water use efficiency more precisely describes specific factors such as plant size, allocation patterns and evaporative demand, which will affect the exchange of carbon dioxide and water through the stomata.</p>

<p>During the last few decades, coral reefs have become a disappearing feature of tropical marine environments, and those reefs that do remain are severely threatened. It is understood that humans have greately altered the environment under which these ecosystems previously have thrived and evoloved. Overharvesting of fish stocks, global warming and pollution are some of the most prominent threats, acting on coral reefs at several spatial and temporal scales. Presently, it is common that coral reefs have been degraded into alternative ecosystem regimes, such as macroalgae-dominated or sea urchin-barren. Although these ecosystems could potentially return to coral dominance in a long-term perspective, when considdering current conditions, it seems likely that they will persist in their degraded states. Thus, recovery of coral reefs cannot be taken for granted on a human timescale. Multiple stressors and disturbances, which are increasingly characteristic of coral reef environments today, are believed to act synergistically and produce ecological surprises. However, current knowledge of effects of compounded disturbances and stress is limited. Based on five papers, this thesis investigates the sublethal response of multiple stressors on coral physiology, as well as the effects of compounded stress and disturbance on coral reef structure and function. Adaptive responses to stress and disturbance in relation to prior experience are highlighted. The thesis further explores how inherent characteristics (traits) of corals and macroalgae may influence regime expression when faced with altered disturbance regimes, in particular overfishing, eutrophication, elevated temperature, and enhanced substrate availability. Finally, possibilities of affecting the resilience of macroalgae-dominaed reefs and shifting the community composition towards a coral-dominated regime are explored.</p>

Bleaching is defined as the disruption of the symbiotic relationship between the host and Symbiodinium dinoflagellates, resulting in the exposure of the calcium carbonate skeleton. This phenomenon has been mostly linked to seawater temperature elevation. Bleaching events are becoming more intense and frequent, harming coral reefs around the world and affecting almost 100% of the community in some places. Therefore, it is important to understand if the effects of these recurrent stress events are cumulative on this photosymbiotic relationship. We experimentally tested how the Symbiodinium-associated jellyfish Cassiopea andromeda is affected by three consecutive thermal stress events, monitoring bell size and chlorophyll-a concentration. Jellyfishes in four physiological conditions regarding feeding frequency (daily and every three days) and Symbiodinium concentration (low and high) were subject to a one-week thermal stress at each experimental round. Three treatments were applied: 27ºC (control, maintained in this temperature during all experiment), 30ºC and 33ºC. After a three-week recovery period at 27ºC, thermal stress events were applied on two more instances intertwined by a recovery period. C. andromeda bleached and its chlorophyll-a content decreased for all temperatures during the first thermal stress and only at 33ºC for the second and third events. Size was only affected by feeding and by the second thermal stress. Higher food offer caused organisms to increase in size while those offered food with lower frequency shrunk. As observed in other hosts, high Symbiodinium concentration resulted in loss of chlorophyll-a, probably due to excessive production of reactive oxygen species and self-shading. C. andromeda jellyfish bleached just in the first stress event and its resilience to multiple thermal stresses is dependent upon biotic and abiotic factors. There was no difference in mortality in any condition or temperature applied. Feeding plays a crucial role on size and symbiotic dependency under thermal stress. Symbiodinium concentration is an important factor in experiments testing climate change scenarios and although symbiont concentration has no relation to the growth of the host, it affects the Symbiodinium itself, thus might be considered in future experiments. Chronic effects of multiple thermal stresses were observed for chlorophyll-a, while these were acute for growth. C. andromeda apparently does not rely only on Symbiodinium, especially under stressful conditions and can be used as a model to investigate the effects of climate change in Symbiodinium symbiosis. Without major host mortality, the behavior of Symbiodinium in its physiological limits after every single bleaching event can be monitored. / O branqueamento é definido pela ruptura da relação simbiótica entre o hospedeiro e o dinoflagelado Symbiodinium, resultando na exposição do esqueleto de carbonato de cálcio do hospedeiro, e tem sido associado principalmente à elevação da temperatura da água do mar. Esses eventos estão se tornando mais intensos e frequentes, prejudicando recifes de corais ao redor do mundo, resultando em branqueamento de quase 100% da comunidade em alguns locais. Portanto, é importante entender se os efeitos desses eventos de estresse são cumulativos neste relacionamento fotossimbiótico. No presente estudo foi experimentalmente testado como medusas de Cassiopea andromeda em simbiose com Symbiodinium são afetadas por três eventos consecutivos de estresse térmico, monitorando o tamanho da umbrella e a concentração de clorofila-a. Medusas em quatro condições fisiológicas em relação à frequência de aporte heterotrófico (diário e a cada três dias) e a concentração de Symbiodinium (baixa e alta) foram sujeitas a um estresse térmico de uma semana em cada rodada experimental. Foram aplicados três tratamentos: 27ºC (controle, mantido nesta temperatura durante todo o experimento), 30ºC e 33ºC. Após um período de recuperação de três semanas a 27ºC, os eventos de estresse térmico foram aplicados em mais duas instâncias interligadas por um período de recuperação. C. andromeda branqueou e sua clorofila-a diminuiu para todas as temperaturas durante o primeiro estresse térmico e apenas a 33ºC para o segundo e terceiro eventos. O tamanho só foi afetado pela alimentação e pelo segundo estresse térmico. A oferta de alimento mais frequente fez com que os organismos aumentassem de tamanho, enquanto aqueles aos quais se oferecia com uma menor frequência, diminuíram de tamanho. Conforme observado em outros hospedeiros, a alta concentração de Symbiodinium resultou em perda de clorofila-a, provavelmente devido à produção excessiva de espécies reativas de oxigênio e auto-sombreamento. Medusas de C. andromeda branquearam apenas no primeiro evento de estresse e sua resiliência a múltiplos estresses térmicos depende de condições bióticas e abióticas. Não houve diferença na mortalidade em qualquer condição ou temperatura aplicada. Sob condições de estresse térmico, o aporte heterotrófico desempenha um papel crucial no tamanho e dependência da simbiose. A concentração de Symbiodinium é um fator importante nos experimentos que utilizam cenários de mudanças climáticas. Embora a concentração de simbiontes não tenha relação com o crescimento do hospedeiro, isso afeta o próprio Symbiodinium e, portanto, deve ser considerado em futuros experimentos. Efeitos crônicos de múltiplos estresses térmicos foram observados para a clorofila-a, enquanto estes são agudos para o crescimento. C. andromeda aparentemente não depende do Symbiodinium, especialmente sob condições estressantes e pode ser usado como modelo para investigar os efeitos das alterações climáticas no Symbiodinium em simbiose. Sem maior mortalidade do hospedeiro, o comportamento do Symbiodinium pode ser monitorado até seus limites fisiológicos após cada evento de branqueamento.

Bleaching is defined as the disruption of the symbiotic relationship between the host and Symbiodinium dinoflagellates, resulting in the exposure of the calcium carbonate skeleton. This phenomenon has been mostly linked to seawater temperature elevation. Bleaching events are becoming more intense and frequent, harming coral reefs around the world and affecting almost 100% of the community in some places. Therefore, it is important to understand if the effects of these recurrent stress events are cumulative on this photosymbiotic relationship. We experimentally tested how the Symbiodinium-associated jellyfish Cassiopea andromeda is affected by three consecutive thermal stress events, monitoring bell size and chlorophyll-a concentration. Jellyfishes in four physiological conditions regarding feeding frequency (daily and every three days) and Symbiodinium concentration (low and high) were subject to a one-week thermal stress at each experimental round. Three treatments were applied: 27ºC (control, maintained in this temperature during all experiment), 30ºC and 33ºC. After a three-week recovery period at 27ºC, thermal stress events were applied on two more instances intertwined by a recovery period. C. andromeda bleached and its chlorophyll-a content decreased for all temperatures during the first thermal stress and only at 33ºC for the second and third events. Size was only affected by feeding and by the second thermal stress. Higher food offer caused organisms to increase in size while those offered food with lower frequency shrunk. As observed in other hosts, high Symbiodinium concentration resulted in loss of chlorophyll-a, probably due to excessive production of reactive oxygen species and self-shading. C. andromeda jellyfish bleached just in the first stress event and its resilience to multiple thermal stresses is dependent upon biotic and abiotic factors. There was no difference in mortality in any condition or temperature applied. Feeding plays a crucial role on size and symbiotic dependency under thermal stress. Symbiodinium concentration is an important factor in experiments testing climate change scenarios and although symbiont concentration has no relation to the growth of the host, it affects the Symbiodinium itself, thus might be considered in future experiments. Chronic effects of multiple thermal stresses were observed for chlorophyll-a, while these were acute for growth. C. andromeda apparently does not rely only on Symbiodinium, especially under stressful conditions and can be used as a model to investigate the effects of climate change in Symbiodinium symbiosis. Without major host mortality, the behavior of Symbiodinium in its physiological limits after every single bleaching event can be monitored. / O branqueamento é definido pela ruptura da relação simbiótica entre o hospedeiro e o dinoflagelado Symbiodinium, resultando na exposição do esqueleto de carbonato de cálcio do hospedeiro, e tem sido associado principalmente à elevação da temperatura da água do mar. Esses eventos estão se tornando mais intensos e frequentes, prejudicando recifes de corais ao redor do mundo, resultando em branqueamento de quase 100% da comunidade em alguns locais. Portanto, é importante entender se os efeitos desses eventos de estresse são cumulativos neste relacionamento fotossimbiótico. No presente estudo foi experimentalmente testado como medusas de Cassiopea andromeda em simbiose com Symbiodinium são afetadas por três eventos consecutivos de estresse térmico, monitorando o tamanho da umbrella e a concentração de clorofila-a. Medusas em quatro condições fisiológicas em relação à frequência de aporte heterotrófico (diário e a cada três dias) e a concentração de Symbiodinium (baixa e alta) foram sujeitas a um estresse térmico de uma semana em cada rodada experimental. Foram aplicados três tratamentos: 27ºC (controle, mantido nesta temperatura durante todo o experimento), 30ºC e 33ºC. Após um período de recuperação de três semanas a 27ºC, os eventos de estresse térmico foram aplicados em mais duas instâncias interligadas por um período de recuperação. C. andromeda branqueou e sua clorofila-a diminuiu para todas as temperaturas durante o primeiro estresse térmico e apenas a 33ºC para o segundo e terceiro eventos. O tamanho só foi afetado pela alimentação e pelo segundo estresse térmico. A oferta de alimento mais frequente fez com que os organismos aumentassem de tamanho, enquanto aqueles aos quais se oferecia com uma menor frequência, diminuíram de tamanho. Conforme observado em outros hospedeiros, a alta concentração de Symbiodinium resultou em perda de clorofila-a, provavelmente devido à produção excessiva de espécies reativas de oxigênio e auto-sombreamento. Medusas de C. andromeda branquearam apenas no primeiro evento de estresse e sua resiliência a múltiplos estresses térmicos depende de condições bióticas e abióticas. Não houve diferença na mortalidade em qualquer condição ou temperatura aplicada. Sob condições de estresse térmico, o aporte heterotrófico desempenha um papel crucial no tamanho e dependência da simbiose. A concentração de Symbiodinium é um fator importante nos experimentos que utilizam cenários de mudanças climáticas. Embora a concentração de simbiontes não tenha relação com o crescimento do hospedeiro, isso afeta o próprio Symbiodinium e, portanto, deve ser considerado em futuros experimentos. Efeitos crônicos de múltiplos estresses térmicos foram observados para a clorofila-a, enquanto estes são agudos para o crescimento. C. andromeda aparentemente não depende do Symbiodinium, especialmente sob condições estressantes e pode ser usado como modelo para investigar os efeitos das alterações climáticas no Symbiodinium em simbiose. Sem maior mortalidade do hospedeiro, o comportamento do Symbiodinium pode ser monitorado até seus limites fisiológicos após cada evento de branqueamento.

Thesis: Ph. D., Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Department of Biology; and the Woods Hole Oceanographic Institution), 2017. / Cataloged from PDF version of thesis. / Includes bibliographical references (pages 151-163). / Population connectivity is a fundamental process that governs the spatial and temporal dynamics of marine ecosystems. For many marine species, population connectivity is driven by dispersal during a planktonic larval phase. The ability to obtain accurate, affordable, and meaningful estimates of larval dispersal patterns is therefore a key aspect of understanding marine ecosystems. Although field observations provide insight into dispersal processes, they do not provide a comprehensive assessment. Individual-based models (IBMs) that couple ocean circulation and particle-tracking models provide a unique ability to examine larval dispersal patterns with high spatial and temporal resolution. Obtaining accurate results with IBMs requires simulating a sufficient number of particles, and the sequential Bayesian procedure presented in chapter 2 identifies when the number of particles is adequate to address predefined research objectives. In addition, this method optimizes the particle release locations to minimize the requisite number of particles. Even after applying this method, the computational expense of IBM studies is still large. The model in chapter 3 seeks to increase the affordability of IBM studies by transferring some of the calculations to graphics processing units. Chapter 4 describes three algorithms that assist in interpreting IBM output by identifying coherent geographic clusters from population connectivity data. The first two algorithms have existed for nearly a decade and recently been applied separately to marine ecology, and we provide a direct comparison of the results from each. Additionally, we develop and present a new algorithm that simultaneously considers multiple species. Finally, in chapter 5, we apply these tools and a trait-based modeling framework to assess which species traits are most likely to impact dispersal success and patterns in the Gulf of Maine. We conclude that the traits influencing spawning distributions and habitat requirements for settlement are most likely to influence dispersal. / by Benjamin Thomas Jones. / Ph. D.

Contaminants may affect marine organisms through various pathways with impacts evident across a variety of spatial and temporal scales. Organisms may encounter short pulsed exposures which contaminate surface waters for hours to days, or more persistent but patchy contamination of benthic habitats throughout their entire life-cycle. This thesis examines the responses of epifauna associated with macroalgae to a pulsed exposure of contaminants (storm-water input) and to chronic contamination via metal accumulation within temperate algal beds. The effects of storm water were monitored during a two-year survey of Sydney Harbour which sampled epifauna before and after heavy rainfall. Epifaunal assemblages declined throughout the harbour following storm events but for the most part these declines were not attributable to storm-water runoff. However, transient (&lt 4 d) and localized impacts of storm water upon physico-chemical characteristics of recipient water and some epifaunal groups were identified around storm drains. A novel field dosing technique tested the relative importance of freshwater and associated metals as causative agents of behavioural avoidance and direct mortality responses. Strong avoidance of storm-water plumes was found which could be entirely explained by freshwater inundation, with no additional effects of metals. No direct mortality was observed following brief exposures. Contaminants introduced by storm water may accumulate within the tissues of macroalgae and potentially pose persistent threats to epifauna. Colonisation of epifauna was reduced on algae with enhanced copper levels, and the nesting behaviour, feeding and survival of an abundant amphipod were all negatively affected by copper load. Subsequent field surveys identified sufficient copper, lead and zinc contamination in Sydney Harbour algal beds to pose direct toxic threats to epifauna. The abundance of herbivorous amphipods correlated negatively with the copper content of a common algal species. However, differences in metal accumulation between algal species resulted in spatially variable levels of contamination. Small-scale patchiness of contaminants within these landscapes may allow populations of mobile species to persist if contaminated hosts are avoided. In summary, epifaunal assemblages appeared resilient to storm-water pulses. Recovery of affected groups was rapid and large fluctuations in abundance appear to be part of the natural flux of epifaunal communities. In contrast, assemblages responded strongly to algal-bound contaminants and this has emerged as an important pathway of contaminant exposure and impact within algal habitats.

<p>Meiofauna, and especially marine nematodes are common in sediments around the world. Despite very wide ranging distributions in many nematode species, little is presently known about their dispersal mechanisms shaping these patterns. Rafting, and perhaps ballast water transport has been suggested as viable means for nematode long-range transport. On a much smaller scale other processes have been suggested for their dispersal. They generally include some form of passive suspension into the water column and later on a passive, haphazard settling back towards the bottom.</p><p>Small-scale phenomena in nematode dispersal were studied by conducting a series of studies at Askö field station, Trosa Archipelago, Baltic proper. Studied aspects were one case of macrofaunal influence on nematode dispersal rate, using an amphipod, <i>Monoporeia affinis</i> as disturbing agent, and three different studies on mechanisms related to settling. The experiments were conducted both in laboratory and field settings.</p><p>The amphipod <i>Monoporeia affinis</i> did not exert any influence on the dispersal rate in the nematodes. The nematode dispersal was only an effect of time, in the aspect that the more time that past, the more nematodes dispersed from their place of origin. The settling experiments revealed that nematodes do have an active component in their settling behaviour, as they were able to exert influence on the spot where they were to settle. They were able to choose settling spot in response to the food quality of the sediment. It also became evident that contrary to common belief, nematodes are able to extend their presence in the water column far beyond the times that would be predicted considering settling velocities and hydrodynamic conditions alone.</p>

<p>The work in this thesis deals with effects of changed species richness on process rates among stream-living macroinvertebrates. Global biodiversity is decreasing rapidly and it is poorly known what the consequences of this loss may be for ecosystems and the services they provide. Hence, it is important to investigate the potential effects of losing species. In streams, deforestation, introduction of non-native species, pollution and channelization are examples of events that may affect species richness negatively. In this thesis emphasis is on changes in species richness within functional feeding groups (FFGs) of stream-living macroinvertebrates. The FFGs used were shredding detritivores, grazers, filter feeders and predators - all of which uphold important ecological processes in streams. Along with an observational field study, species richness was manipulated in laboratory and field experiments to investigate the effects of changed species richness on process rates and thus ecosystem functioning.</p><p>The results show that effects of changed species richness on process rates may be dramatic. Among the shredding detritivores there were negative effects on leaf mass loss, regardless whether fixed, random or predicted sequences of species loss was investigated. These effects could be attributed to either species richness per se or species composition. However, among the other FFGs the relationship between species richness and process rates was less consistent. In filter feeders, there was no or a negative effect of decreasing species richness while both grazers and predators showed positive effects of species loss.</p><p>The results also show that the most important interactions between species in an experiment, thus potentially in a natural community, are likely to determine what the effect of species loss on process rates will be. Facilitation and niche differentiation lead to reduced process rates if species are lost, while mechanisms, such as interspecific resource or interference competition, produce the opposite effect. Furthermore, in systems with a diminishing resource, the first two mechanisms may become more important over time enhancing the effect of species loss in the long term.</p><p>In conclusion, effects of species loss may be dramatically negative or positive even if lost species are classified as redundant. The effect in the short term most likely depends on which species are lost, on the original species composition and on the underlying mechanisms. Questions remaining to be answered are how important the observed effects are in more complex systems and if they are persistent over time? Future studies will tell. </p>

<p>This thesis deals with the dynamics of tundra living voles with emphasis on the most common one, the grey-sided vole (Clethrionomys rufocanus). The tundra area chosen for the study was Finnmarksvidda, a vast flatland in northernmost Norway. All small mammal herbivores in the area showed dramatic fluctuations, and field experiment were conducted in order to elucidate these density fluctuations. The specific subjects addressed included: 1/ Temporal and spatial appearance of density fluctuations of voles and lemmings in the area, 2/ The generality of the density patterns observed, 3/ The impact of predation by vole predators during summertime, 4/ The impact of grey-sided vole grazing on food plants of different preference in a predator free environment, in the presence and absence of extra food, and 5/ The impact of food availability on density and demography of grey-sided voles in a predator free environment.</p><p>The results achieved showed that voles in the slope and lowland had cyclic density fluctuations with 5 years duration. The cycles consisted of four phases: an increase phase, a peak phase, a decline phase and a crash phase. In the unproductive lowland and on the moderately productive slope, small pockets of productive habitats seemed to work as “triggers” for the cycles. The lemming fluctuations in the upper plateau (separated from the slope by a steep zone of boulders) differed markedly from the vole patterns in the lowland.</p><p>Only two lemming peaks were recorded in twenty years. Both peaks had very short increase phases, a knife-sharppeak phase and no decline phase before the crash. A comparison between our results and lemming and vole populations from two other areas in Fennoscandia revealed the same difference in fluctuation pattern between lemmings and voles as seen in our area. This results suggests that lemmings in barren tundra highlands and voles in slightly more productive tundra lowlands are regulated by different mechanisms.</p><p>The exclusion of vole predators from vole populations during summertime led to increase in overall vole density. Densities of the clumsy field vole (Microtus agrestis) and juveniles of all species showed the strongest positive effects of the exclusion.</p><p>An experiment analysing the effects of food availability was conducted in islands in a large lake where grey-sided voles were introduced to predator free islands . Supplemental food was given to the voles in two unproductive, and two productive islands. Two unproductive and two productive islands were used as reference islands. The density of voles and the vole weight were higher in both the islands with supplemental food and those with high natural productivity. Increased vole density did not significantly increase grazing damage to plants. The cyclic density pattern of the voles in the nearby mainland (that harboured resident vole specialist predators as stoat and weasel) showed little resemblance to the seasonal fluctuations found in the islands (devoid of resident vole specialist predators). This result suggested that predation by stoat and weasel on grey-sided vole populations may cause the cyclic vole fluctuations seen in the area.</p>